'¥.6675 
Vol. 73, pp. 67-82 10 August I960 
PROCEEDINGS 
OF THE 
BIOLOGICAL SOCIETY OF WASHINGTON 
VARIATION IN THE CUBAN LIZARD 
LEIOCEPHALUS RAVICEPS COPE 
By Albert Schwartz 
Department of Biology, Albright College, Reading, Pa. 
Of the five species of the genus Leioeephalus inhabiting 
Cuba, the least known is Leioeephalus ravieeps Cope. De- 
scribed by Cope in 1862 on the basis of specimens collected 
by the botanist Charles Wright in "eastern Cuba," L. ravieeps 
was unknown to Gundlach except from the original description 
(Gundlach, 1875:354; 1880:34). Barbour (1914:301) followed 
Boulenger in regarding L. ravieeps as a synonym of L. vittatus 
(= L. cuhensis), but later he and Ramsden (1919:173) fol- 
lowed Stejneger (1917:53) in affirming the distinctness of the 
species, although they were not certain that it was Cuban. 
These two authors assumed that Wright's types came most 
probably from the Sierra de Yateras ( probably owing to Gund- 
lach's (1880) comment), but Ramsden was unable to find the 
lizard in that immediate area. This is not surprising, when it 
is known that L. ravieeps is an inhabitant of the most xeric areas 
in Cuba, and is not known to occur in mesic areas or forest. 
Cochran (1934:39) apparently was the first to report L. 
ravieeps from a definite locality in Cuba; she cited specimens 
from four localities in the vicinity of Guantanamo Bay in Ori- 
ente, and erroneously (see Schwartz, 1959:110-11) from the 
Doce Leguas keys off Camaguey Province. Alayo (1951:109; 
L955:16) reported the species from La Socapa, on the* west 
side of the Bahia de Santiago, and from Laguna de Baconao 
on the coast road between Santiago de Cuba and Guantanamo. 
Thus, /.. ravieeps is now known from the region of the Bahia 
de Santiago east to the Bahia de Guantanamo; the onl) record 
of the species to the west of the Bahia de Santiago is Alayo's 
specimen From La Socapa, whereas the only record From east 
12 — Prcx . Biol. So< . w ish.. Vol. 73. I960 I 87 I 
^H*^ ^ 
9 
■NSTmmoN AUG lo i960 
i ■ 
68 Proceedings of the Biological Society of Washington 
of the Bahia de Guantanamo is that of Cochran from Boqueron. 
During the summer of 1959, specimens of L. raviceps were 
collected at various localities on the south Oriente coast by 
myself, Ronald F. Klinikowski, and Barton L. Smith. During 
the Christmas holiday season of 1959, a new series was taken 
in the Sierra de Purial by myself and James R. Talada. All 
these collections were made under a National Science Founda- 
tion research grant. For the first time, ample series of speci- 
mens from various localities are available. It is not surprising 
that two subspecies are readily distinguishable in the material 
from Oriente. In addition to these fresh specimens, I have had 
the opportunity to examine the cotypes of L. raviceps in the 
U. S. National Museum, as well as a small series collected by 
Thomas M. Uzzell, Jr., and Richard Etheridge at the U. S. 
Naval Base on the east side of Guantanamo Bay, and a single 
specimen from Baracoa. I wish to thank Doris M. Cochran, 
Norman E. Hartweg, and Miguel L. Jaume Garcia for permis- 
sion to examine specimens respectively in the U. S. National 
Museum (USNM), Museum of Zoology, University of Michi- 
gan (UMMZ), and the Museo y Biblioteca de Zoologia de la 
Habana ( MBZH ) . Knowing my interest in the species, Messrs. 
Uzzell and Etheridge kindly allowed me to examine their very 
pertinent material. The illustrations are the work of Ronald 
F. Klinikowski; his work on my behalf is sponsored by a Na- 
tional Science Undergraduate Research Participation grant. 
In addition to the specimens from Oriente, we collected a 
series of L. raviceps on the northern coast of Matanzas Province 
near Varadero. These specimens are the first extra-Oriente 
lizards of this species, and they too represent a distinct new 
form; a distance of some 690 kilometers separates this popula- 
tion from the nearest known Oriente population of L. raviceps. 
In order to establish which of the three populations of L. 
raviceps best agrees with Cope's concept of the species, I have 
examined the seven cotypes of the species ( USNM 4162 ) . This 
series consists of two adult males and five adult females, all 
in fair condition considering their preservation for almost a 
century. Cope's (op. cit. :183) description of coloration and 
pattern are a composite of the series; he stated, "Above yellow- 
ish brown, with many short, narrow, black longitudinal lines, 
which are something arranged as a double series of dorsal 
Variation In the Cuban Lizard 69 
spots." In general, the males have the dorsal short lines 
(dashes), whereas the females have a double series of dorsal 
spots. This characterization of the cotypes, along with Cope's 
comment "Top of head light yellowish brown," is sufficient to 
restrict it to the population of L. raviceps which occurs east of 
the Bahfa de Guantanamo, i.e., between Guantanamo and 
Cajobabo, and in the Sierra de Pnrial. The remainder of Cope's 
description is consonant with the allocation of the cotypes with 
this population, and examination of the series of cotypes con- 
firms this arrangement. Comparison of the types with fresh 
material from this region, and with specimens of the new dark- 
race from farther west along the Oriente coast, shows that this 
area is occupied by the nominate form. 
Only Gundlach (1880:34) has attempted to restrict the type 
locality of L. raviceps Cope; this Cuban herpetologist restricted 
it to "the mountains near Guantanamo, Oriente." This general 
locality was not unlikely, since it is known that Wright col- 
lected in this area. Repeated search for raviceps in the mesic 
Sierra de Yateras by herpetologists yielded no specimens. How- 
ever, the species does occur in the very different and xeric Si- 
erra de la Vela to the southeast of Guantanamo on the east 
of the Bahfa de Guantanamo. Thus if Gundlach's type locality 
restriction of "the mountains near Guantanamo" is understood 
to refer to the Sierra de la Vela or other of the dry coastal 
ranges, this locality may be regarded as correct. 
I have seen no specimens from Guantanamo itself; I suspect 
that raviceps occurs to the south of that city, on the west side 
of the Bahfa, in the xeric and Opuntia-studded plains which 
occur close to the city itself. We collected one specimen near 
Caimanera. To the southeast of Guantanamo, between the city 
and the U. S. Naval Base, no raviceps were collected, and in 
general this area now, at least, appears unsuitable. However, 
as one approaches the Naval Base, the typical xeric features 
of the flora and landscape appeal - , and the Species should oc- 
cur there, as it does in the Sierra de la Vela. 
The population of L. raviceps between the Bahfa de Santiago 
and the Bahfa de Cuantanamo differs iu coloration, pattern, 
and certain Features of scutellation From that to the east; For 
this western population I propose the name iu honor of Thomas 
70 Proceedings of the Biological Society of Washington 
M. Uzzell, Jr., who collected the first fresh specimens of L. 
raviceps which I had seen, as: 
Leiocephalus raviceps uzzelli, new subspecies 
Type: American Museum of Natural History (AMNH) 79321, from 
18.2 kilometers east of Siboney, Oriente Province, Cuba, taken 25 July 
1959; one of a series obtained by Ronald F. Klinikowski, Albert Schwartz, 
and Barton L. Smith. Original number 7867. See Fig. 1. 
Paratypes: AMNH 79310-20, 79322-35, same data as type. 
Specimens examined and not designated as paratypes: AMNH 79336- 
40, 26.6 km E Siboney; AMNH 79341-45, Laguna de Baconao, 21.8 mi 
E Siboney; AMNH 79346, 2 mi N Caimanera. 
Distribution: The southern Oriente coast from the Bahia de Santiago to 
the Bahia de Guantanamo. 
Diagnosis: A subspecies of L. raviceps characterized by dark tan colora- 
tion with sharply differentiated darker brown lateral fields, relatively 
prominent dark brown to black dorsal dashes or paired blotches, usually 
well defined postorbital blotch and more often incomplete than complete 
supraorbital semicircles. 
Description of type: An adult male, with the following measurements 
(in millimeters) : snout-vent length 65; tail 96, complete; snout to anterior 
border of tympanic opening 13.7; head width 11.4; supraocular scales 
7/7; loreals 3; temporals 9; enlarged auricular scales 4/4; median head 
scales 4; prefrontal row complete 3 scales; frontoparietal row complete 
5 scales; parietals in contact; semicircles incomplete; dorsal crest scales 
occiput to vent 65; dorsal crest scales occiput to axilla 27; scales around 
half body at midbody 33; fourth toe subdigital tricarinate scales 25/26. 
Coloration: The dorsal coloration in life is yellowish tan. There is a 
darker brown lateral field, which in life contained scattered yellow and 
red scales, extending from the eye to the groin. The dorsal surface of the 
head is likewise yellowish tan with some of the scales, especially the inner 
margins of the semicircles, outlined by black. The dorsum between the 
lateral fields shows a median zone which is set off laterally by a longi- 
tudinal paler zone. Ventral to the lateral field there is a longitudinal pale 
whitish band which extends from above the tympanic opening to the groin; 
this line is outlined below by a faintly gray longitudinal band, which 
merges gradually into the pale gray venter. The dorsal crest scales are 
not set off by their coloration from the remainder of the median dorsal 
tan band. The median zone includes a series of about 13 more or less 
triangular dark brown blotches, their apices pointed posteromedially; the 
anterior blotches are darker and more rectangular than the posterior ones. 
There are a few short, dark longitudinal dashes between the blotches. The 
next adjacent longitudinal zones, which have a paler ground color, are 
marked with scattered longitudinal dashes, involving two to three scales. 
The lateral fields are marked with diagonal dark dashes, which are prom- 
inent on the sides and somewhat fainter on the neck; these dashes con- 
tinue more ventrally between the limbs to midway between the limb in- 
Variation hi the Cuban Lizard 
71 
Fig. 1-3 
sertions. Scattered light scales in and below the lateral fields were yellow 
in life; these tight Scales continue onto tlie abdomen as a series of trans- 
\erse rows of white dots against the pale gray of the abdomen. The dorsal 
Surfaces of the limbs are tan; the hindlimbs ha\ e a combination of short 
72 Proceedings of the Biological Society of Washington 
brown dashes and scattered cream colored scales on their dorsal surfaces, 
and these markings continue onto the dorsal surface of the foot. The post- 
orbital blotch is represented by a slightly gray area on the cheek, with a 
few brownish scales intermingled with the gray. The upper labials are 
suffused with tan; the lower labials have deeper tan pigment along the 
infralabial sutures, giving the lower lip a somewhat mottled appearance. 
Three white bands radiate ventrally from the eye, the posterior two sep- 
arated by a vertical black bar which extends ventrally from the middle 
of the eye. The entire ventral surface is pale gray, and the throat is im- 
maculate. The underside of the hindlimbs and the posterior third of the 
abdomen have scattered brown dots; these dots extend as well onto the 
basal portion and sides of the tail. In addition, cream colored scales occur 
on the underside of the hindlimbs and tail. Thus these members are pale 
gray with scattered brown spots, and more abundant cream colored spots. 
Variation: In snout-vent length, 21 adult male L. r. uzzelli average 
63.2 mm (55-71); 9 adult females average 48.1 (45-53). Dorsal crest 
scales (combined data for both sexes) in occiput-vent length average 
63.4 (56-69), and dorsal crest scales in occiput-axilla average 26.0 (23- 
31 ) . One half scales at midbody average 32.4 ( 29-36 ) , loreals 3.3 ( 2-5 ) ; 
temporals, 11.0 (8-13), subdigital fourth toe tricarinate scales 24.3 (22- 
28 ) . The parietals are always in contact, and the supraorbital semicircles 
are more often incomplete (53%) than not. The supraoculars are most 
often 7/7 (68%), with 6/6 occurring as the next most frequent (13%) 
category. Variation in this character includes in addition counts of 5/5 
(3%), 5/6 (6%), 6/7 (6%), and 7/8 (3%). If all counts which in- 
clude at least 7 supraoculars (at least unilaterally) are combined, 77% 
of the population is differentiated from the 23% which have counts 
ranging from 5/5 to 6/6. 
Since there is a certain amount of sexual dimorphism in pattern, the 
two sexes will be discussed separately. Males can be distinguished from 
females in that the former have two pairs of enlarged postanal scales. All 
males are relatively darkly pigmented in life, with a conspicuous lateral 
field as compared with the paler r. raviceps. Some males demonstrate a 
yellowish suffusion of the head scales in life, but in general this has dis- 
appeared in preservative; the more common condition in uzzelli is the 
unicolored tan head scales. The dorsal pattern is like that described for 
the type in most cases — a series of paired dark rectangular or triangular 
blotches in the median tan zone. Some specimens lack these blotches 
and have them supplanted by a uniform series of longitudinal dashes or 
even dark dots; any combination of these patterns may occur on the same 
individual. The lateral fields are always dark and prominent with bold 
black diagonal dashes. The postorbital blotch varies in intensity, but the 
type is unusual in the faint pigmentation of the postorbital area; usually 
the cheek is covered by a heavy black blotch, or there is at least some black 
pigment in this area, and the "blotch" may appear as a hollow or open- 
ended rectangle, brown to tan centrally; the blotch is, even at its most 
reduced, usually better defined and more prominent than it is in r. ravi- 
Variation In the Cuban Lizard 
73 
Fig. 4-10 
ceps. Unregenerated tails have a pattern oi about IT dorsal chevrons, 
their apices pointed posteriorly; the ground color of the tail is faintly pink- 
ish in life, and this serves to distinguish these lizards in the field. The 
throat usually is immaculate, varying in color From white to very pale 
lavender to gray, With occasional clear cream colored scales scattered on 
its margins. Some males show a few gray lines or dots on the throat; this 
gray pigmentation occurs in juveniles more regularly than adults, but it 
appears not to be strictly ontogenetic in nature. The undersides of the 
hindlimbs, posterior abdomen, and basal portion of the tail are dotted w itb 
dark brown as are these members in the type. 
'I he females present the same dark appearance as the males, bul are 
74 Proceedings of the Biological Society of Washington 
more distinctly lined longitudinally (cf. Schwartz, op. cit., sexual di- 
morphism in L. cubensis and L. stictigaster) . The female pattern is an 
intensification of that described for the males — the paired dorsal blotches 
(only one female shows any additional dorsal pigmentation in the form 
of dark dashes in the median zone). The sides have the diagonal dark 
dashes which may be somewhat inconspicuous due to the dark coloration 
of the lateral fields. The hindlimbs have dorsal dark dashes, but these 
may be obscure. The postorbital blotch is usually demonstrated by a 
slightly darker brown area on the cheek. The females, in contrast to the 
males, almost always have dark gray dots, often aligned into longitudinal 
rows, on the throat, and usually lack any dark dots on the underside of the 
hindlimbs, although the belly usually has brown dots on its posterior 
third. Both sexes have the pale facial markings described for the type, but 
they are more clearly expressed in the females than the males. Usually 
there are but two, rather than three, light subocular areas, the anteriormost 
being obscured by the deposition of tan pigment, and the posterior two 
being separated by a definite vertical black bar. The juvenile males re- 
semble the females in pattern and pigmentation very closely. 
All specimens but six have four median head scales (see Fig. 4); five 
specimens have five median head scales ( see Fig. 5 ) and one has six ( see 
Fig. 6). This condition of five and six, rather than four, scales is caused 
either by the unequal transverse division of one of a pair of scutes or by 
the unequal division of both members of a pair. 
Comparisons: L. r. uzzelli differs from its relative, L. r. raviceps, to the 
east in both coloration and scalation, as follows: Although both races are 
tan dorsally, uzzelli is by far the darker of the two; raviceps has a distinctly 
faded or "washed out" appearance ( see Fig. 2 ) , whereas uzzelli is darker. 
Male uzzelli have the lateral fields very dark, and the dorsum is marked 
either with dark brown or black dashes, often organized into two series 
of paired black or brown blotches on either side of the dorsal midline. 
This feature seems to be a retention into the adult of the juvenile (and 
female) blotched pattern. Even if the dorsal pattern is composed of 
dashes, there are at least two pairs of blotches in the nuchal area, and thus 
at times the subspecies uzzelli may superficially resemble L. m. macropus. 
The postorbital blotch is usually demonstrated, in contrast to the condi- 
tion in raviceps, and when best expressed, is black and outlined anteriorly 
by a light line extending ventrally from the eye onto the posterior supra- 
labials, and extends posteriorly to the tympanic opening. Ventrally, males 
of the two subspecies are much alike, except that uzzelli males often have 
a few gray lines or dots on the throat, and the undersurface of the hind- 
limbs, basal portion of the tail and posterior abdomen are much more 
heavily and profusely marked with dark brown dots. In contrast to the 
dull face and labial markings of raviceps, uzzelli has distinct facial mark- 
ings of dark brown or black consisting of three lines (one just below the 
canthus rostralis, another from the anterior corner of the eye, and the third 
from the posterior half of the eye) which encloses three light areas and 
extend onto the supralabials. The infralabials are regularly mottled white 
Variation In the Cuban Lizard 75 
and dark brown to tan. Female uzzclli resemble female raviceps except 
that the latter females are much paler and appear faded, and lack the 
female uzzclli face pattern of a single vertical black line through the cen- 
ter of the eye. The gray throat markings of female uzzclli are more pro- 
nounced and darker than those of raviceps, and one female demonstrates 
brown dots on the underside of the hindlimbs. 
As noted above, the two races differ in general coloration, raviceps 
being more distinctly yellowish-tan than uzzclli. The yellowish suffusion 
on the head of raviceps is not shown so distinctly in uzzclli, although oc- 
casional specimens do show it. L. r. uzzclli shows yellow or red dots (lat- 
eral scales) on the sides and in the area of the shoulder, and apparently 
lacks the green lateral scales of raviceps. 
Males of the two subspecies do not differ in snout-vent length; 23 adult 
male raviceps average 66.5 mm (61-71). Eleven adult females average 
54.5 (52-57); it is possible that the females of nzzelli are consistently 
smaller than those of raviceps, but there are too few uzzelli females avail- 
able to be certain. At present the females of the two races can almost be 
separated by size alone (largest female uzzelli 53 mm, smallest female 
raviceps 52 mm). Means and extremes of scale characters of raviceps 
are: dorsal crest scales in occiput-vent length 64.2 (55-74); dorsal crest 
scales in occiput-axilla length 27.5 (22-32); one half scales at midbody 
32.3 (27-36); loreals 3.6 (2-5); temporals 11.6 (9-15). None of these 
counts are significantly different from those of uzzelli. However, tricari- 
nate subdigital fourth toe scales 25.5 (21-30) in raviceps; inspection of 
Fig. 11 indicates that this count is significantly higher than that of uzzclli. 
Both races usually have four median head scales; only one r. raviceps ex- 
amined had five median head scales. Both races usually have 7/7 supra- 
oculars; 71% of r. raviceps have 7 or 8 supraoculars at least unilaterally. 
Parietal contact occurs in 95% of the r. raviceps; the semicircles are com- 
plete in 81% of the lizards. This is in contrast to uzzelli, in which 47% of 
the specimens have complete semicircles; the difference is significant. 
Remarks: The south Oriente coast lies in the rain shadow of the Sierra 
Maestran system; specifically, between the Bahia de Santiago and the 
Bahia de Guantanamo, the range involved (which is considered as part 
of the Sierra Maestra by Marrero, 1951: 584) is the Cordillera de la Gran 
Piedra. North and east of the Guantanamo Basin are the Sierra del Guaso, 
the Sierra de Maquey and the Siena de la Vela. The extensive Sierra de 
Purial (Las Cuchillas) limits the very narrow coastal area from east of 
Guantanamo to Cabo Maisi. Marrero (op. cit.i 658) described the coast 
from Guantanamo to Cabo Maisi in the following accurate terms: "The 
southern coast of Oriente, from Guantanamo eastward, is in reality a des- 
ert, with infrequent rains during the entire year and a \erophvtie \egeta- 
tion. in which cacti predominate, interrupted only by the gallery vege- 
tation at the base of palms, together with the rare rivers which descend 
to the ocean." Later, Marrero (p. 859) mentioned the botanist Victorin's 
appellation for pari of this region as the "inferno of Maisi"; such descrip- 
tions are verbally adequate to describe the intense he.it, sandy and rock} 
76 Proceedings of the Biological Society of Washington 
soils, and xerophytic plants which characterize this coastal strip. The 
coast from Santiago to Guantanamo, while nonetheless hot and xeric, is 
less so than the Guantanamo area, and supports occasional stands of de- 
ciduous trees, mangroves, and palms, along with some low herbaceous 
cover. L. raviceps is widespread between Guantanamo and Cabo Maisi, 
but appears to be less so and more restricted to the niche of sandy soil and 
Opuntia between Santiago and Guantanamo. However, in this latter re- 
gion, it has been found sparingly in other habitats, which are invariably 
dry. For example, at Laguna de Baconao, we took both r. uzzelli and 
m. macropus in close proximity to one another, at the edge of the man- 
grove border of the lake. However, macropus occurred within the shady 
forest on mud and moist ground, whereas uzzelli shunned this cooler 
habitat in preference to the dry and extremely hot hillsides. 
Although the single specimen of L. raviceps from Baracoa (MBZH 136) 
is the only individual from the north coast of Oriente, and is separated 
from the south coast r. raviceps population by the Sierra de Purial, this 
lizard does not differ in any way from the southern coastal specimens of 
r. raviceps. It has the herringbone dorsal pattern which occurs in south 
coast r. raviceps and not in uzzelli. Although we collected Leiocephalus 
in the Baracoa area, no raviceps were taken. The series of lizards from 
the Sierra de Purial (AMNH 83791-806, from 4.6 mi N Cajobabo) shows 
that in extreme eastern Cuba the distribution of this lizard may not be 
completely coastal. Under proper conditions it may invade non-coastal 
montane areas, provided that the soil and moisture conditions are suit- 
able. At this Cajobabo locality, the lizards were taken along the Cajo- 
babo-Baracoa road in a dry and sunny road cut, as they ran about on the 
broken shale fragments where there was little or no plant cover. Such 
relatively isolated populations between the south coast and Baracoa may 
well maintain genetic continuity between the populations of the two coasts. 
On the other hand, r. raviceps may continue to Baracoa via a more or 
less continuous population from the south coast around Cabo Maisi and 
thence to Baracoa. 
The absence of L. raviceps west of the Bahia de Santiago is puzzling; 
as noted previously, the only record for the species in this area is at La 
Socapa, which is on the west side of the bay itself. The common Leio- 
cephalus on the coast south of the Sierra Maestra is L. macropus; there 
are no obvious differences in the vegetation or substrate of this coastal 
strip as compared to that of the Santiago-Guantanamo strip except the 
absence of the sand-Opuntia niche. It is possible that raviceps and mac- 
ropus are competitors, and that the former has been unable to become 
established to the west of Santiago; another possibility is that the western 
coast is too moist for this species and the apparent absence of sandy soils 
and Opuntia is a limiting factor. 
The completely unexpected occurrence of L. raviceps in Matanzas 
Province is indeed surprising. The series of 22 lizards from two localities 
near Varadero is very different in several features from L. raviceps in 
Variation In the Cuban Lizard 77 
Oriente, and accordingly, in honor of Ronald F. Klinikowski, the dis- 
coverer of this western population, I propose the name: 
Leiocephalns raviceps klinikowskii, new subspecies 
Type: AMNH 83326, adult male, from 4.5 kilometers southwest of 
Varadero, Mantanzas Province, Cuba, taken 8 September 1959, one of 
a series collected by Ronald F. Klinikowski, Albert Schwartz, and Barton 
L. Smith. Original No. 8528. See Fig. 3. 
Paratypes: AMNH 83327-46, same data as type; AMNH 83347, 5.5 
kilometers southwest of Varadero, Mantanzas, 8 September 1959, R. F. 
Klinikowski. 
Distribution: Known only from the coast southwest of Varadero in 
Matanzas Province, Cuba. 
Diagnosis: A subspecies of L. raviceps characterized by very pale yel- 
lowish tan dorsal coloration, lateral fields very inconspicuous and hardly 
differentiated from ground color, dorsum with prominent black dorsal 
dashes, poorly defined postorbital spot, usually 5 or 6 median head scales 
and 6/6 supraoculars, and high number of dorsal crest scales and scales 
around midbody. 
Description of type: An adult male, with the following measurements 
in mm and counts: snout-vent length 69; tail 74, distal one third regen- 
erated; snout to anterior border of tympanic opening 14.4; head width 
11.8; supraocular scales 6/6; loreals 5; temporals 16; enlarged auricular 
scales 4/4; median head scales 6; prefrontal row complete 3 scales; fronto- 
parietal row complete 5 scales; parietals in contact; semicircles complete; 
dorsal crest scales occiput to vent 62; dorsal crest scales occiput to axilla 
27; scales around one half body at midbody 35; fourth toe subdigital tri- 
earinate scales 24/26. 
Coloration: The dorsal coloration in life is pale yellowish-tan, almost 
sand colored, with the dorsal crest scales immaculate pale yellow. There 
is little evidence of longitudinal zonation and the lateral fields are not 
appreciably darker than the dorsum itself, nor are they separated from 
the ventral gray coloration by an additional band of tan; the effect lat- 
erally is a gradual blending of the dorsal yellowish-tan coloration into the 
gray of the abdomen. The dorsal surface of the head is yellowish-tan with 
black pigment stippled over the head scutes, especially the prefrontals, 
posterior Supraoculars, and the parietals and interparietal. The entire 
dorsum and lateral field area is marked with short black longitudinally 
aligned dashes (see Fig. 3), which become obscure on the neck and form 
two very faint pairs of nuchal blotches which are barely discernible. 
Cream Colored isolated scales on the sides extend onto the abdomen in a 
series of five parallel rows, with additional cream scales on the abdomen 
anterior to the hindh'inbs. Some of the t.m scales in the lateral field area 
were Orange basally in lite. The Ion-limbs are mottled tan dorsally and 
the finuers are immaculate pale tan, giving the lizard a distinctly pale 
fingered appearance. The hindlimbs show short black longitudinal dashes 
on all sections, hut they are arranged as spots on the loot itself; there are 
78 Proceedings of the Biological Society of Washington 
additional light scales on the dorsal surface of the hindlimbs. The post- 
orbital blotch is represented by a vertical black bar behind the eye, and a 
darker irregular blotch posterior to this bar; the cheek itself is tan. Three 
gray lines radiate from the eye, one below the canthus rostralis, another 
from the anterior corner of the eye, and a third from the center of the 
eye; these extend to the supralabials, and are continued on the infralabial 
sutures. The entire ventral surface is gray, with a darker gray suffused 
area across the chin, another on the throat, and a few scattered cream 
scales on the throat. The undersurface of the hindlimbs is almost white, 
again with a few light tan dots scattered over the shank; darker dots oc- 
cur on the posterior abdomen and on the underside of the tail. The un- 
regenerated portion of the tail shows about six dull gray chevrons with 
the apices directed posteriorly; the ground color of the tail is dull reddish- 
tan in life. 
Variation: In snout-vent length, four adult male L. r. klinikowskii aver- 
age 58.5 mm (53-69); eight adult females average 51.9 mm (46-59). 
Dorsal crest scales ( combined data for both sexes ) in occiput-vent length 
average 66.2 (61-71), and dorsal crest scales in occiput-axilla length 
average 27.0 (24-29). One half scales at midbody average 35.3 (32-39), 
loreals 4.2 (4-5), temporals 13.9 (11-16), subdigital fourth toe tricari- 
nate scales 25.9 (21-27). The parietals are more often (86%) in con- 
tact than not, and the supraorbital semicircles are always complete. 
Of the three paratypic males, one resembles the type in dorsal pattern; 
the remaining two individuals have the dorsal zonation somewhat more 
apparent, and still retain some remnants of the juvenile male paired blotch 
pattern. All have pale yellowish-tan dorsa, very pale fingers, immaculate 
yellow dorsal crest scales, chevrons (about 18) on the complete tails, 
rather diffuse gray postorbital blotches, black stippling on the head scutes, 
and dark facial and labial markings. The three paratypic males lack brown 
dots on the undersurface of the hindlimbs, but show them on the under- 
side of the base of the tail. One has a completely immaculate throat, and 
the remaining two have a gray suffusion and scattered diffuse gray dots. 
The eight adult females are much as the females of raviceps and uzzelli 
except that they are much paler; all but one show the paired female 
blotches and dashes in the lateral fields. The single exception is unusually 
gray and has dashes in the lateral field area and very pale gray dashes 
dorsally. All females have black on the head scutes, postorbital blotches 
absent or if present diffuse and gray, and a contrasting face pattern of 
gray or black lines. The inter-infralabial sutures are distinctly black, and 
the throat always has some indication of gray dots or dashes. The under- 
side of the hindlimbs is usually immaculate white, but a few individuals 
have indications of very light tan dots. The dark brown belly dots are 
present and rather extensive. 
There are 10 juvenile female paratypes ranging in snout-vent lengths 
from 26 to 44 mm. These show the same pattern as the adult females and 
differ only in the more definite dark gray chin and throat markings. 
Nineteen of the 22 specimens of klinikowskii have five or six median 
Variation In the Cuban Lizard 79 
head scales; variation in this feature is shown in Figs. 4 to 10. Only two 
individuals have four median head scales (Figs. 4 and 7). Fig. 7 shows 
that, although there are only four head scales in this case, anterior con- 
tinuation of the incipient median suture would convert this pattern to five 
median scales. The other lizard with four median head scales differs in 
no wise from four-head-scaled individuals of raviceps and uzzelli ( Fig. 4 ) . 
Thus almost 100% of klinikowskii have either 5 or 6 median head scales, 
at least incipiently. 
Variation in number of supraoculars is great; 6/6 occurs in the most 
individuals (12 lizards, 55%), and 6/7 in six lizards (27%). Only two 
(9%) klinikowskii have 7/7 supraoculars, in contrast to 68% in uzzelli 
and 44% in raviceps. Other counts in klinikowskii are 6/8 and 7/8 (one 
individual each). Counts of more than 6/6 arise in the following three 
ways: ( 1 ) a supraocular, usually the sixth, is divided longitudinally so that 
two scales occupy the area of one scale; (2) the sixth is replaced by two 
scales transversely, thus giving the usual 7 supraoculars of raviceps and 
uzzelli; (3) a small supraocular is added anteriorly from fusion or enlarge- 
ment of one of the semicircle scales. The occurrence of six supraoculars 
at least unilaterally is shown in all but one specimen (which has 7/8) of 
klinikowskii. 
Comparisons: L. r. klinikowskii differs from both raviceps and uzzelli 
in its very pale coloration, light to no ventral spotting on the hindlimbs, 
usually very inconspicuous lateral fields, relatively faint postorbital blotch, 
pale fingers, and immaculate yellow dorsal crest scales. In scalation, the 
Matanzas race differs in usually 6/6 supraoculars, although a relatively 
high percentage (32%) of raviceps has this number of supraocular scales. 
From uzzelli, klinikowskii differs in having the semicircles always com- 
plete, whereas the former has the semicircles more often incomplete. In- 
spection of Fig. 11 shows that klinikowskii differs significantly from uzzelli 
in number of dorsal crest scales in occiput-vent length, from raviceps and 
uzzelli in number of one half midbody scales, and from uzzelli in number 
of fourth toe scales. 
In size, klinikowskii males average smaller than males of both the other 
races; this is probably an artifact of the small size of the sample of male 
klinikowskii. Female klinikowskii reach a larger size than females of the 
other races, although the average of the series is intermediate between 
that of raviceps and uzzelli. It is especially interesting that, despite the 
apparent great hiatus between the known range of klinikowskii and uzzelli, 
the scale differences between these two races are not more striking. 
Remarks: The habitat of L. r. klinikowskii is apparently the sandy 
beaches of Punta de Hicacos. It is probable that the race has a wider 
distribution than presently known; since much of the beach area of the 
VaraderO area is now converted to recreational use, collecting must be 
carried on at less populated areas. The type locality is a stretch of open 
beach, with almost no herbaceous cover except Ij)<>mca and scattered tufts 
of t^rass, separated from the Matanzas— Varadero highway (Via Blanca) 
l>y a stand of introduced Casuaiina. The single lizard From 5.5 kilometers 
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Variation In tlxc Cuban Lizard 81 
southwest of Varadero was taken on sandy soil in rather dense grassy 
cover. The lizards were all captured by hand; this was a relatively simple 
task since there was such little cover. They sought refuge in land-crab 
burrows, and when especially hard pressed, would bury themselves in 
the sand much in the same manner of Uma and would quickly disappear 
from view. 
I suspect that other populations of L. raviceps remain to be discovered. 
Sandy areas on the north coast from Matanzas to northern Oriente may 
well harbor disjunct populations of this lizard. The only other comparable 
area which I have visited on the north coast is Playa Santa Lucia in east- 
ern Camaguey; no raviceps were collected there. However, it is interest- 
ing that the type locality of L. /'. klinikoicskii was visited in the summer 
of 1958, and these lizards were not observed at this or any other collecting 
locality along the Punta de Hicacos' north coast. Visits in 1958 and 1959 
to a locality 13 kilometers northeast of Matanzas did not yield specimens 
of L. raviceps, and I doubt that the species occurs in this immediate re- 
gion; here there is no beach as such, the shore having a formation of diente 
de perro limestone with sandy dunes behind the limestone, covered with 
dense herbaceous and woody vegetation which is apparently unsuitable 
for L, r. klinikoicskii. 
Literature Cited 
Alayo Dalmau, Pastor. 1951. Especies herpetologicas halladas en San- 
tiago de Cuba. Bol. Hist. Nat. Soc. Felipe Poey, 2(7): 106- 
110. 
. 1955. Lista de los reptiles de Cuba. Museo Charles T. 
Ramsden, Santiago de Cuba, pp. 1-29, 7 pis. 
Barbour, Thomas. 1914. A contribution to the zoogeography of the 
West Indies, with especial reference to amphibians and rep- 
tiles. Mem. Mus. Comp. Zool., 44(2): 209-359, 1 pi. 
and Charles T. Ramsden. 1919. The herpetology of Cuba. 
Mem. Mus. Comp. Zool., 47(2): 71-213, 15 pis. 
Cochran, Doris M. 1934. Herpetological collections from the West 
Indies made by Dr. Paul Bartsch under the Walter Rathbone 
Bacon Scholarship, 1928-30. Smith. Misc. Coll., 92(7): 1-48. 
Cope, E. D. 1862. Contributions to neotropical saurology. Proc. Acad. 
Nat. Sci. Philadelphia, pp. 176-188. 
Gundlach, Juan. 1875. Catalogo de los reptiles cubanos. Anal. Soc. 
Espan. Hist. Nat., 4: 347-368. 
. 1880. Contribucion a la erpetologia eubana. G. Montiel y 
Cia., La Habaua, pp. 1-98. 
Marrero, Levi. 1951. Geografia de Cuba. Alfa, La Ilabana, pp. i-\\i, 
1-736, 478 figs. 
Schwartz, Albert. 1959. Variation in lizards of the LeiocephaltlS cu- 
beruis complex in Cuba and [sla de Pinos. Bull. Fla. State 
Mus., 4(4): 97-143, L0 figs. 
82 Proceedings of the Biological Society of Washington 
Stejneger, Leonhard. 1917. Cuban amphibians and reptiles collected 
for the United States National Museum from 1899 to 1902. 
Proc. U. S. Nat. Mus., 53: 259-291, 128 figs. 
Explanation of Figures 
Fig. 1. — Leiocephalus r. uzzelli, new subspecies, dorsal view of type 
(AMNH 79321). 
Fig. 2. — Leiocephalus r. raviceps, dorsal view (AMNH 79350). 
Fig. 3. — Leiocephalus r. klinikowskii, new subspecies, dorsal view of 
type (AMNH 83326). 
Figs. 4 to 10. — Dorsal view of median head scales of L. raviceps show- 
ing variation in number from four to six scales; figures from specimens as 
follows: 4. AMNH 79314; 5. AMNH 83327; 6. AMNH 79310; 7. AMNH 
83328; 8. AMNH 83331; 9. AMNH 83333; 10. AMNH 83326. 
Fig. 11. — Counts of dorsal crest scales in occiput-vent length, scales 
around one half of body at midbody, and subdigital tricarinate scales on 
fourth toe in three Cuban subspecies of Leiocephalus raviceps. Horizontal 
line indicates range of variation in sample; vertical line, the mean; hollow 
rectangles indicate two standard errors on each side of the mean. If rec- 
tangles on two sets of data do not overlap, a statistically significant differ- 
ence between the means is suggested. Sizes of individual samples are 
given to the left of each line. Higher number of individual counts in 
fourth toe scales is due to counting the number of subdigital scales on the 
fourth toe of both the right and left foot.